Disassembling Papaver: a restriction site analysis of chloroplast DNA

The results from a chloroplast DNA restriction site analysis of the genera Papaver and Roemeria of subf. Papaveroideae (Papaveraceae) and five outgroup taxa are presented. Papaver is represented by 14 species of eight of the 11 sections recognized, Roemeria by two of its three species. Hunnemannia fumariifolia (subf. Eschscholzioideae), Chelidonium majus (subf. Chelidonioideae), Romneya coulteri, Argemone munita and Stylomecon heterophyllum (all subf. Papaveroideae) were chosen as outgroups. DNAs were digested with 24 restriction enzymes. The major results from this analysis are: 1. Papaver, Roemeria and Stylomecon form a monophyletic group supported by at least 17 restriction site mutation…

Phylogeny and quaternary history of the European montane/alpine endemicSoldanella(Primulaceae) based on ITS and AFLP variation

Soldanella contains 16 species of herbaceous perennials that are endemic to the central and south European high mountains. The genus is ecogeographically subdivided into forest/montane and alpine species. Evolutionary relationships and large-scale biogeographic patterns were inferred from parsimony analyses of the internal transcribed spacer (ITS) regions of nuclear ribosomal DNA, and genetic distance analyses based on amplified fragment length polymorphism (AFLP) markers. The ITS region proved useful for examining subgeneric relationships and testing hypotheses on genus-wide divergence times, whereas the AFLP markers were suitable for studying relationships among closely related taxa and b…

Morphological evolution and ecological diversification of the forest-dwelling poppies (Papaveraceae: Chelidonioideae) as deduced from a molecular phylogeny of the ITS region

Sequences of the ITS region of nrDNA were analyzed for the seven genera of Papaveraceae subf. Chelidonioideae s.str. Three major clades can be recognized. These are 1.Chelidonium/Hylomecon/Stylophorum, 2.Eomecon/Sanguinaria, and 3.Bocconia/Macleaya. The monophyly of genera in the first of these three clades is doubtful, and clades two and three are sister to each other. Use of the ITS phylogeny of the subfamily to trace its morphological and ecological evolution shows that morphological change is concentrated in theBocconia/Macleaya clade, and probably related to the evolution of wind-pollination from insect-pollination in these two genera after habitat shift.

Western Eurasian-western North American disjunct plant taxa: The dry-adapted ends of formerly widespread north temperate mesic lineages-and examples of long-distance dispersal

The phylogeny of the Papaveraceae sensu lato: morphological, geographical and ecological implications

On the background of the phylogeny of the Papaveraceae s.l., the evolution of the androecium and gynoecium, the geographical distribution, and the ecological differentiation of the family is discussed. An effort is made to homologize the diverse androecia of Pteridophyllum, Hypecoum, Fumariaceae and Papaveraceae s.str., and it is concluded that both the polyandrous androecium as found in the Papaveraceae s.str. and the polycarpellate gynoecium as found in Papaveraceae s. str. subfam. Papaveroideae incl. Platystemonoideae are secondary and derived from whorled oligomerous androecia and bicarpellate gynoecia, respectively. The comparative consideration of the geographical distribution, ecolog…

Asterales: Introduction and Conspectus

Long-distance seed dispersal, clone longevity and lack of phylogeographical structure in the European distributional range of the coastal Calystegia soldanella (L.) R. Br. (Convolvulaceae)

Aim  To explore the relative effects of Quaternary climatic history vs. species-specific biological properties (high seed dispersability, high seed longevity, clonal growth) on phylogeographical structure in European Mediterranean and Atlantic Ocean and Black Sea material of the coastal dune plant Calystegia soldanella (L.) R. Br. Location  Black Sea and European Mediterranean and Atlantic Ocean coasts. Methods  Variation in amplified fragment length polymorphism was analysed at two different sampling levels. First, an entire-range sample from the Black Sea to the North Sea, including single individuals from sites evenly spread along this entire coast was analysed. Second, in a population-l…

The effect of Quaternary climatic changes on plant distribution and evolution

Abstract Climatic oscillations in the Quaternary have played a major role in changing the geographical distribution of plant species. Recent molecular work has provided new insights into the location of glacial refugia and routes of postglacial expansion. A comparative analysis shows that phylogeographic patterns in Europe appear to be less congruent than in North America. The change of geographical distribution has also provided opportunities for speciation through isolation, and hybridization after secondary contact. However, a complete understanding of the significance of the Quaternary climatic changes for plant speciation requires further research into the estimation of reliable and ac…

Phylogeography ofSyringa josikaea(Oleaceae): Early Pleistocene divergence from East Asian relatives and survival in small populations in the Carpathians

Tertiary relict plant species of Europe have had a large distribution range before the Pleistocene but today are confined to small refugial areas. Syringa josikaea of the largely East Asian genus Syringa is a shrub of temperate forests in the Carpathians, restricted to altogether 25 small populations in two disjunct areas, the Apuseni Mountains (Romania) and the Ukrainian Carpathians. Miocene and Pleistocene fossil remains indicate the long-term presence of the species in Central Europe; hence S. josikaea has been considered a Tertiary relict. We aimed at clarifying the historical biogeography of S. josikaea by estimating the divergence time between S. josikaea and its Asian relatives, and …

Asian relationships of the flora of the European Alps

Background: It has long been recognised that the Alps have strong biogeographical links to Asia. Aims: We investigate here whether the two areas are connected either through southern (mountain chains between the Alps and the Himalayas) or through northern regions. Methods: We compiled the geographical distribution of plant genera growing both in the Alps and the Himalayas, and outside these two regions. This compilation was used to search for molecular phylogenetic literature which was screened for area relationships informative for our purposes. Results: Of 933 genera growing in the Alps, 653 were considered further. Of the 429 genera growing in the Alps and the Himalayas, 218 grow in both…

Understanding Mediterranean‐Californian disjunctions: molecular evidence from Chenopodiaceae‐Betoideae

Chenopodiaceae subfam. Betoideae is distributed in both western Eurasia (four genera) and western North America (one genus). To understand the origin of this disjunction, the phylogeny of the subfamily was reconstructed and dated using ndhF, matK/trnK, tmL-trnF spacer, and ITS sequence variation, penalized likelihood and Langley-Fitch, and calibration with three different fossils. Maximum Parsimony and Maximum Likelihood analyses of the molecular data show that Betoideae are monophyletic, but that relationships of the Himalayan Acroglochin, traditionally included in Betoideae because of the shared possession of a circumscissile capsule, are uncertain. Among the betoidean genera, Beta (excl.…

Dual colonization of the Palaearctic from different regions in the Afrotropics bySenecio

Aim Investigation of the geographical origin of Senecio and the colonization of the Palaearctic including the relationship of life-history strategy and elevational distribution in the source area and the colonized area. Location Worldwide with a focus on the Afrotropic and the Palaearctic. Methods Sampling focused on adding species from the Afrotropic to existing datasets of Senecio. Two nuclear markers and three chloroplast markers were amplified and sequenced. Maximum likelihood and Bayesian inference were used to infer phylogeny, divergence times, biogeographical history and life-history strategy evolution. Results Senecio originated most likely during the Mid to Late Miocene in western …

The Palaeoanthropocene – The beginnings of anthropogenic environmental change

Abstract As efforts to recognize the Anthropocene as a new epoch of geological time are mounting, the controversial debate about the time of its beginning continues. Here, we suggest the term Palaeoanthropocene for the period between the first, barely recognizable, anthropogenic environmental changes and the industrial revolution when anthropogenically induced changes of climate, land use and biodiversity began to increase very rapidly. The concept of the Palaeoanthropocene recognizes that humans are an integral part of the Earth system rather than merely an external forcing factor. The delineation of the beginning of the Palaeoanthropocene will require an increase in the understanding and …

What is the origin of the Scottish populations of the European endemic Cherleria sedoides (Caryophyllaceae)?

Cherleria sedoides L. (Minuartia sedoides (L.) Hiern) is a montane perennial which, with some species in Minuartia sect. Spectabiles, is more closely related to Scleranthus than to other Minuartia species and is therefore best restored to the reinstated and redefined genus Cherleria. Reconstruction of the ancestral area of the clade containing C. sedoides suggests that it evolved in the Alps or the Balkan peninsula. The species now has an unusual distribution, being present in the mountains of southern Europe and Scotland but absent from the Arctic. Three historical scenarios that might have led to the presence of the species in Scotland are outlined and tested by a molecular analysis compa…

Patterns and causes of incongruence between plastid and nuclear Senecioneae (Asteraceae) phylogenies

One of the longstanding questions in phylogenetic systematics is how to address incongruence among phylogenies obtained from multiple markers and how to determine the causes. This study presents a detailed analysis of incongruent patterns between plastid and ITS/ETS phylogenies of Tribe Senecioneae (Asteraceae). This approach revealed widespread and strongly supported incongruence, which complicates conclusions about evolutionary relationships at all taxonomic levels. The patterns of incongruence that were resolved suggest that incomplete lineage sorting (ILS) and/or ancient hybridization are the most likely explanations. These phenomena are, however, extremely difficult to distinguish beca…

Genetic variation of naturally growing olive trees in Israel: from abandoned groves to feral and wild?

Background Naturally growing populations of olive trees are found in the Mediterranean garrigue and maquis in Israel. Here, we used the Simple Sequence Repeat (SSR) genetic marker technique to investigate whether these represent wild var. sylvestris. Leaf samples were collected from a total of 205 trees at six sites of naturally growing olive populations in Israel. The genetic analysis included a multi-locus lineage (MLL) analysis, Rousset’s genetic distances, Fst values, private alleles, other diversity values and a Structure analysis. The analyses also included scions and suckers of old cultivated olive trees, for which the dominance of one clone in scions (MLL1) and a second in suckers (…

An ITS phylogeny of tribe Senecioneae (Asteraceae) and a new delimitation of Senecio L.

Senecioneae is the largest tribe ofAsteraceae, comprised ofca. 150 genera and 3,000 species. Approximately one-third of its species are placed in Senecio, making it one of the largest genera of flowering plants. Despite considerable efforts to classify and understand the striking morphological diversity in Senecioneae, little is known about its intergeneric relationships. This lack ofphylogenetic understanding is predominantly caused by conflicting clues from morphological characters, the large size ofthe tribe, and the absence of a good delimitation of Senecio. Phylogenetic analyses of nrITS and plastid DNA sequence data were used to produce a hypothesis of evolutionary relationships in Se…

Phylogeny and evolution of the Arctium-Cousinia complex (Compositae, Cardueae-Carduinae)

The phylogeny and evolution of the Arctium-Cousinia complex, including Arctium, Cousinia as one of the largest genera of Asteraceae, Hypacanthium and Schmalhausenia, is investigated. This group of genera has its highest diversity in the Irano-Turanian region and the mountains of Central Asia. We generated ITS and rpS4-trnT-trnL sequences for altogether 138 species, including 129 (of ca. 600) species of Cousinia. As found in previous analyses, Cousinia is not monophyletic. Instead, Cousinia subgg. Cynaroides and Hypacanthodes with together ca. 30 species are more closely related to Arctium, Hypacanthium and Schmalhausenia (Arctioid clade) than to subg. Cousinia (Cousinioid clade). The Arctio…

A revision of Gynura (Asteraceae: Senecioneae)

A revision of the paleotropical genus Gynura Cass. is presented. Forty-four species are recognized, of which three are described as new. These are G. daviesiae Vanijajiva & Kadereit, G. villosa Vanijajiva & Kadereit, and G. siamensis Vanijajiva & Kadereit. Gynura dissecta (F. G. Davies) Vanijajiva & Kadereit, G. annua (F. G. Davies) Vanijajiva & Kadereit, and G. aurantiaca (Bl.) DC. subsp. parviflora (F. G. Davies) Vanijajiva & Kadereit are new combinations. An identification key, descriptions, and distribution maps are provided.

Ecological differentiation and reproductive isolation of two closely related sympatric species of Oenanthe (Apiaceae)

Oenanthe conioides is a lower Elbe endemic plant species growing in the freshwater tidal zone around Hamburg (Germany). Its closest relative Oenanthe aquatica is widely distributed in Eurasia and grows in calm and shallow freshwater. The two species differ in habitat requirements but are otherwise sympatrically distributed, suggesting that ecological divergence has to be maintained in the face of gene flow. In the present study, we investigated ecological differentiation and reproductive isolation in these two species. An amplified fragment length polymorphism analysis found clear genetic differentiation between the two species implying reproductive isolation. A reciprocal transplantation e…

Genetic evidence for divergent selection onOenanthe conioidesandOe. aquatica(Apiaceae), a candidate case for sympatric speciation

The opportunity for habitat shift in sympatry is thought to be an important factor in sympatric speciation by facilitating assortative mating and offering opportunities for divergent selection. Oenanthe conioides (Apiaceae) is a narrow endemic from the lower Elbe river area (Germany) where it is restricted to areas experiencing fresh water tides inundating the plants twice a day. The species was shown to have originated from Oe. aquatica which is widely distributed in Europe and grows in still or slowly flowing fresh water. Reciprocal transplant experiments have previously shown that in both habitats the non-native species is less fit than the native, and several phenotypic traits have been…

Phylogeny and Biogeography of <I>Epimedium</I>/<I>Vancouveria</I> (Berberidaceae): Western North American - East Asian Disjunctions, the Origin of European Mountain Plant Taxa, and East Asian Species Diversity

Using ITS and atpB-rbcL spacer sequences of 38 (of 55) species of the highly disjunct Eurasian/North African Epimedium and all three species of its western North American sister genus Vancouveria, we reconstructed the phylogeny of these two genera and dated major splits with a molecular clock approach. Epimedium was found to be monophyletic with a stem age dated to between 9.7 and 7.4 million years ago (My). Within Epimedium, almost all sections as recognized in the most recent classification of the genus were found to be monophyletic but subg. Epimedium was found to be paraphyletic in relation to subg. Rhizophyllum. Range formation in Eurasia proceeded as follows: in a first step, the west…

Range-wide phylogeography of the European temperate-montane herbaceous plantMeum athamanticumJacq.: evidence for periglacial persistence

Aim The aim of this study is to analyse the genetic population structure of Meum athamanticum Jacq. in order to explore the alternative hypotheses (1) that the central and northern highland populations are the result of post-glacial recolonization from southern refugia, and the disjunct distribution of M. athamanticum can be explained by modern ecological conditions, or (2) that extant populations north of the Alps and Pyrenees persisted in situ during glacial periods. Location Europe. Methods Variation of amplified fragment length polymorphisms (AFLPs) was analysed for 23 populations from the entire range of the species. We used band-based approaches and methods based on allele frequencies…

The role of hybridization in the evolution ofCousinias.str.(Asteraceae, Cardueae)

Abstract Mehregan I. & Kadereit J. W.: The role of hybridization in the evolution of Cousinia s.str. (Asteraceae, Cardueae). — Willdenowia 39: 35–47. — Online ISSN 1868-6397; © 2009 BGBM Berlin-Dahlem. doi:10.3372/wi.39.39102 (available via http://dx.doi.org/) In order to assess the possible role of interspecific hybridization for the evolution of the high diversity of Cousinia subg. Cousinia (Cousinia s.str., c. 600 species), we examined the frequency of hybridization in this taxon. For this, hypothetical hybrid combinations published in the literature (28 putative hybrids and 11 intermediate forms) were critically examined. In addition, two hybrids were identified in the present study by …

Integrating species distribution models (SDMs) and phylogeography for two species of AlpinePrimula

The major intention of the present study was to investigate whether an approach combining the use of niche-based palaeodistribution modeling and phylo-geography would support or modify hypotheses about the Quaternary distributional history derived from phylogeographic methods alone. Our study system comprised two closely related species of Alpine Primula. We used species distribution models based on the extant distribution of the species and last glacial maximum (LGM) climate models to predict the distribution of the two species during the LGM. Phylogeographic data were generated using amplified fragment length polymorphisms (AFLPs). In Primula hirsuta, models of past distribution and phylo…

Identity and relationships of Sempervivum tectorum (Crassulaceae) in the Rhine Gorge area

Sempervivum tectorum (Crassulaceae), an orophyte widespread in the European high mountains, also grows in rocky habitats of the Rhine Gorge area (Upper Middle Rhine, Mosel and Ahr river valleys). On the background of its long history of cultivation, it is unclear whether S. tectorum is native or naturalized in the Rhine Gorge area. Using 52 accessions of S. tectorum from across its geographical range (except SE Europe) as well as 15 samples of S. calcareum and S. marmoreum in our final sample, we conducted a genotyping-by-sequencing analysis. The genetic data were used for the identification of genetic groups and for the reconstruction of phylogenetic relationships. We found that the materi…

The role of in situ species diversification for the evolution of high vascular plant species diversity in the European Alps—A review and interpretation of phylogenetic studies of the endemic flora of the Alps

Abstract By harbouring ca. 3500 native vascular plant species in an area of ca. 170 000 km2, the European Alps represent a region of very high species diversity. Using the most recently published flora of the area and phylogenetic literature, I here review which proportion of the endemic flora of the Alps is the result of in situ diversification, i.e., of diversification in the area and (largely) restricted to the area. There exist only very few and mostly species-poor species diversifications in the Alps, accounting for ca. 1.2% of the native and for ca. 9% of the endemic flora of the Alps. In contrast to this, ca. 33% of the endemic species of the Alps belong to lineages widespread in the…

Phylogeny, biogeography and evolution of Triglochin L. (Juncaginaceae) – Morphological diversification is linked to habitat shifts rather than to genetic diversification

A species-level phylogeny is presented for Triglochin, the largest genus of Juncaginaceae (Alismatales) comprising about 30 species of annual and perennial herbs. Triglochin has an almost cosmopolitan distribution with Australia as centre of species diversity. Trans-Atlantic and trans-African disjunctions exist in the genus. Phylogenetic analyses were conducted based on molecular data obtained from nuclear (ITS, internal transcribed spacer) and chloroplast sequence data (psbA-trnH spacer, matK gene). Based on the phylogeny of the group divergence times were estimated and ancestral distribution areas reconstructed. Our data confirm the monophyly of Triglochin and resolve relationships betwee…

C 3 –C 4 intermediates may be of hybrid origin – a reminder

Summary The currently favoured model of the evolution of C4 photosynthesis relies heavily on the interpretation of the broad phenotypic range of naturally growing C3–C4 intermediates as proxies for evolutionary intermediate steps. On the other hand, C3–C4 intermediates had earlier been interpreted as hybrids or hybrid derivates. By first comparing experimentally generated with naturally growing C3–C4 intermediates, and second summarising either direct or circumstantial evidence for hybridisation in lineages comprising C3, C4 and C3–C4 intermediates, we conclude that a possible hybrid origin of C3–C4 intermediates deserves careful examination. While we acknowledge that the current model of C…

Introduction to the Orders and Families of Uncertain Placement of this Volume

The present volume of this book series is the first of two to (almost) complete the treatment of the Asterids which started with Vol. VI (Cornales, Ericales, 2004), Vol. VII (Lamiales, 2004) and Vol. VIII (Asterales, 2007). It contains the orders Boraginales, Garryales and Solanales of the Lamiids (Asterids I) as well as three unplaced families of that clade, i.e. Vahliaceae, Icacinaceae and Metteniusaceae, and the orders Aquifoliales, Escalloniales, Bruniales, Dipsacales and Paracryphiales of the Campanulids (Asterids II).

Nomenclature ofSoldanellaL. (Primulaceae)

Following a revision of Soldanella, we here synonymize 109 non-hybrid names of this genus. We also lectotypify or neotypify 17 names of Soldanella. These are S. alpina L., S. alpina F. W. Schmidt, S. alpina L. var. carestiae Cristofolini et Pignatti, S. alpina L. f. major Cristofolini et Pignatti, S. austriaca Vierh., S. carpatica Vierh., S. marmarossiensis Kldst., S. minima Hoppe, S. minima Hoppe subsp. samnitica Cristofolini et Pignatti, S. minima Hoppe f. latifolia Cristofolini et Pignatti, S. montana Mikan, S. montana Willd., S. montana Willd. subsp. faceta Kress, S. occidentalis Vierh., S. pusilla Baumg., S. pusilla Baumg. f. calcicola Vierh., and S. pusilla Baumg. var. chrysosplenifol…

Phylogeny of prickly poppies,Argemone (Papaveraceae), and the evolution of morphological and alkaloid characters based on ITS nrDNA sequence variation

Evolutionary relationships withinArgemone (Papaveraceae) were inferred from nucleotide variation in nuclear ribosomal DNA. A complete representation of the genus was achieved by using herbarium material to a large extent (74%). Four distinct clades can be recognized based on the molecular results. The support for the different clades varies greatly due to a very uneven distribution of characters. Although some clades are largely unresolved, some unexpected relationships, for example the sister group relationship of the relict speciesA. subintegrifolia to the rest ofArgemone, were found. The evolution of morphological and alkaloid characters against the background of the obtained phylogeny i…

Late Quaternary distributional stasis in the submediterranean mountain plant Anthyllis montana L. (Fabaceae) inferred from ITS sequences and amplified fragment length polymorphism markers.

Anthyllis montana is a submediterranean, herbaceous plant of the southern and central European mountains. The internal transcribed spacer (ITS) regions of nuclear ribosomal DNA were sequenced from multiple accessions of the species and several closely related taxa. In addition, amplified fragment length polymorphism (AFLP) was analysed from 71 individuals of A. montana collected in 20 localities, mainly in the Pyrenees, Alps, Italian Peninsula and Balkans. Our ITS phylogeny showed a sequential branching pattern in A. montana, implying a western Mediterranean origin followed by an eastward migration. ITS clock calibrations suggest that speciation of A. montana took place at the Pliocene-Plei…

The genetic ghost of an invasion past: colonization and extinction revealed by historical hybridization inSenecio

Hybridization is an important evolutionary factor in the diversification of many plant and animal species. Of particular interest is that historical hybridization resulting in the origin of new species or introgressants has occurred between species now geographically separated by great distances. Here, we report that Senecio massaicus, a tetraploid species native to Morocco and the Canary Islands, contains genetic material of two distinct, geographically separated lineages: a Mediterranean lineage and a mainly southern African lineage. A time-calibrated internal transcribed spacer phylogeny indicates that the hybridization event took place up to 6.18 Ma. Because the southern African lineage…

The systematics of Soldanella (Primulaceae) based on morphological and molecular (ITS, AFLPs) evidence

Based on the observation of more than 6300 herbarium specimens and molecular (ITS, AFLPs) evidence, Soldanella is divided into 16 species and four subspecies. Soldanella angusta, S. oreodoxa and S. rugosa are described as new, and the circumscriptions of S. marmarossiensis, S. major, S. pusilla ssp. pusilla and S. hungarica are emended. Nine hybrid names are lectotypified. All non-hybrid taxa are described in detail and distribution maps are provided. Interspecific relationships are discussed on the background of morphological and molecular variation.

Rapid radiation of North American desert genera of the Papaveraceae: Evidence from restriction site mapping of PCR-amplified chloroplast DNA fragments

Phylogenetic relationships of a group of North American desert genera of the Papaveraceae subfam. Papaveroideae and Platystemonoideae were investigated with an RFLP analysis of three PCR-amplified chloroplast genome regions. In agreement with earlier results it was found that subfam. Platystemonoideae is nested within subfam. Papaveroideae. The group under study is characterized by a large number of generic autapomorphies but only few informative synapomorphies. This is interpreted as strong evidence for a rapid radiation event caused by major climatic changes in the past. Poor phylogenetic resolution seems to reflect biological reality and not to be an experimental artifact. There is also …

Phylogeography of a Tertiary relict plant,Meconopsis cambrica(Papaveraceae), implies the existence of northern refugia for a temperate herb

The perennial herb Meconopsis cambrica, a western European endemic, is the only European species of the otherwise Himalayan genus Meconopsis and has been interpreted as a Tertiary relict species. Using rbcL and ITS sequence variation, we date the split between M. cambrica and its sister clade Papaver s.str. to the Middle to Upper Miocene (12.8 Myr, 6.4–19.2 Myr HPD). Within M. cambrica, cpDNA sequence variation reveals the existence of two groups of populations with a comparable level of genetic variation: a northern group from Great Britain, the Massif Central, the western Pyrenees and the Iberian System, and a southern group from the central and eastern Pyrenees. Populations from the Cant…

RAPD evidence for a sister group relationship of the presumed progenitor-derivative species pairSenecio nebrodensis andS. viscosus (Asteraceae)

The phylogenetic and phenetic analysis of 109 RAPD polymorphisms inS. nebrodensis, a perennial and self-incompatible endemic of four mountain ranges in Spain, andS. viscosus, a self-compatible annual widespread in Europe, as well asS. lividus, S. sylvaticus andS. vulgaris revealed a sister group relationship between the first two species. This result contrasts sharply with the earlier hypothesis based on isozyme variation thatS. viscosus originated from within a paraphyleticS. nebrodensis and that the two species represent a progenitor-derivative pair. After considering possible reasons for the sister group relationship found, including the possibility of rooting artefacts, it is concluded …

The Phylogeny of Linum and Linaceae Subfamily Linoideae, with Implications for Their Systematics, Biogeography, and Evolution of Heterostyly

The genus Linum consists of over 180 species, the most famous being L. usitatissimum, the source of linen and linseed oil. The eight genera of Linaceae subf. Linoideae, of which Linum is the largest, exhibit a complex biogeographic distribution, inhabiting all continents except Antarctica. Numerous species in Linoideae are heterostylous, but the ancestral breeding system of the group has not been determined. We present phylogenetic analyses of 44 species representing all eight genera of subf. Linoideae and 37 species of Linum, with data from the chloroplast (ndhF, trnL-F, trnK3′ intron) and the nuclear ITS, with Hugonia (Linaceae subf. Hugonioideae) as outgroup. Sequences of rbcL from 48 sp…

Introduction to the Orders of this Volume

The present volume of this book series completes the treatment of the Asterids. Asterids are now contained in Vols. VI (Cornales, Ericales, 2004), VII (Lamiales, 2004), VIII (Asterales, 2007), XIV (Aquifoliales, Boraginales, Bruniales, Dipsacales, Escalloniales, Garryales, Paracryphiales, Solanales, Icacinaceae, Metteniusaceae, Vahliaceae, 2016) and the present volume, which contains the orders Apiales and Gentianales (except Rubiaceae). The only families of Asterids not treated in the series are Acanthaceae (Lamiales), Convolvulaceae (Solanales) and Rubiaceae (Gentianales).

Simultaneous speciation in the European high mountain flowering plant genus Facchinia (Minuartia s.l., Caryophyllaceae) revealed by genotyping-by-sequencing

Understanding the relative importance of different mechanisms of speciation in a given lineage requires fully resolved interspecific relationships. Using Facchinia, a genus of seven species centred in the European Alps, we explore whether the polytomy found by Sanger sequencing analyses of standard nuclear (ITS) and plastid markers (trnQ-rps16) is a hard or soft polytomy by substantially increasing the amount of DNA sequence data, generated by genotyping-by-sequencing. In comparison to 142 phylogenetically informative sites in the Sanger sequences the GBS sequences yielded 3363 phylogenetically informative sites after exclusion of apparently oversaturated SNPs. Maximum parsimony, maximum li…

Past, present and future of mountain species of the French Massif Central - the case ofSoldanella alpinaL. subsp.alpina(Primulaceae) and a review of other plant and animal studies

Aim  Our goals were: (1) to investigate patterns of genetic variation in the French Massif Central (MC) of Soldanella alpina (Primulaceae), an alpine plant species that has only one known population in the region; (2) to analyse these patterns in order to deduce the Quaternary history of the population and to predict how current climatic warming may affect it; and (3) to review molecular analyses from the MC to evaluate the importance of the region for the conservation of genetic diversity. Location  Europe, with a special focus on the French Massif Central and adjacent regions. Methods  Amplified fragment length polymorphisms (AFLPs) were analysed for 192 individuals (nine populations) of …

Chloroplast DNA evidence for introgression and long distance dispersal in the desert annualSenecio flavus (Asteraceae)

Phylogenetic analysis of chloroplast DNA (cpDNA) restriction site variation supports a close genetic relationship between the Southwest AsianSenecio flavus subsp.breviflorus and the North AmericanS. mohavensis. The intercontinental disjunct distribution of these two desert annuals may have originated via long distance dispersal. The chloroplast genomes of the Southern and North AfricanS. flavus subsp.flavus and subsp.breviflorus differ by at least ten restriction sites, while at most two restriction sites differentiate the cpDNA genomes of subsp.breviflorus and the outgroupS. squalidus. This suggests that the cpDNA genome ofS. flavus subsp.breviflorus may have resulted from introgression an…

Systematics and Phylogeny

Funktionelle Morphologie und Anatomie der Gefäßpflanzen

In den vorangehenden Kapiteln wurden Zellen als elementare Lebenseinheiten und Bauelemente von Geweben behandelt. Auch im Vielzeller ist jede einzelne Zelle tatsachlich eine elementare Lebenseinheit, aber sie reprasentiert hier nicht den Organismus, dessen makroskopische Gestaltung vom zellularen Bau genauso unabhangig ist wie die Architektur eines Bauwerks aus Ziegeln und anderen Bauelementen. Man kann (und konnte ja tatsachlich lange Zeit) sinnvoll Morphologie betreiben, ohne von Zellen etwas zu wissen. Komplexe Gestaltbildungen sind auch ohne Zellengliederung moglich (Abb. 3.1, Abb. 24.48 D, Abb. 24.76; s. Abschn. 1.2.3.6). Freilich zeigen die Seltenheit echter Groszeller und die enorme …

Tetraploid European Salicornia species are best interpreted as ecotypes of multiple origin

Abstract Salicornia procumbens and S. stricta are two tetraploid European salt marsh species of locally adjacent but ecologically differentiated distribution. Whereas S. procumbens grows in the lowest part of the salt marsh, it is replaced by S. stricta in the middle part (and diploid Salicornias in the upper part). Using AFLPs and a reciprocal transplantation experiment, we investigated whether the two species represent distinct evolutionary lineages. The analysis of AFLP variation clearly showed that both species are not monophyletic. Also, accessions do not cluster according to geographical origin. The transplantation experiment revealed that S. procumbens shows significantly reduced fit…

Historical biogeography in a linear system: genetic variation of Sea Rocket (Cakile maritima) and Sea Holly (Eryngium maritimum) along European coasts

The exclusively coastal Cakile maritima and Eryngium maritimum represent a linear biogeographical system. Genetic variation among 25 individuals of C. maritima and 16 individuals of E. maritimum, from the coasts of Europe, North Africa and the Canary Islands, was analysed using random amplified polymorphic DNAs (RAPDs) and intersimple sequence repeats (ISSRs). Genetic distances (Dice) were calculated and used to investigate the correlation between genetic and geographical distances, to construct Neighbour Joining (NJ) trees, and to compare mean genetic distances between areas within and across species. Genetic distances and geographical distances measured along the coast are well correlated…

Molecular phylogeny of the Old World representatives of Papaveraceae subfamily Papaveroideae with special emphasis on the genus Meconopsis

The RFLP-analysis of PCR amplified cpDNA fragments of 42 representatives of Papaveraceae subfam. Papaveroideae resulted in six most parsimonious cladograms. The subfamily can be divided into a New World group (Arctomecon, Argentone, Canbya, Romneya and Platystemon) and an Old World group (Meconopsis, Papaver s.l. and Roemeria) containing Stylomecon heterophylla and Papaver californicum as New World taxa. In the Old World group neither Meconopsis nor Papaver are monophyletic. Whereas Meconopsis consists of three clades, Papaver comprises five clades, with Roemeria as sister group to P. sect. Argemonidium and Stylomecon as sister group to P. californicum. Various lines of evidence suggest tha…

The comparative biology of the closely related Senecio nebrodensis and S. viscosus, a narrow endemic and a widespread ruderal

The closely related Senecio nebrodensis, a perennial herb from four mountain ranges in Spain, and S. viscosus, an annual herb widespread in Europe, are being compared by the study of herbarium material and comparative cultivation. In cultivation, S. nebrodensis behaved as an annual, completing its life cycle within a shorter period of time than S. viscosus. The formation of basal leaf rosettes is much more pronounced in S. nebrodensis than in S.viscosus. The main difference between the two species, however, is their breeding system. While S. nebrodensis is self-incompatible, S. viscosus is self-compatible. Differences in reproductive morphology (mainly size and number of ray florets) are su…

Late Quaternary history ofHippophaë rhamnoidesL. (Elaeagnaceae) inferred from chalcone synthase intron (Chsi) sequences and chloroplast DNA variation

Fossil pollen records indicate that Hippophae rhamnoides (Sea Buckthorn) was widespread on late- and early postglacial raw soils throughout much of central and northern Europe, but that Early Holocene reforestation restricted populations to northern coastal habitats, or along mountain streams in the Alps, Pyrenees, and Carpathians. We used sequence variation at the nuclear chalcone synthase intron ( Chsi ), in conjunction with chloroplast DNA–restriction fragment length polymorphism data, to investigate the intraspecific phylogeny, phylogeographic structure, and expansion demographic history of this dioecious and wind-pollinated shrub at its range-wide scale in Europe and Asia Minor. Four m…

<I>The Genus</I> Meconopsis. <I>Blue Poppies and Their Relatives</I> (2014) by C. Grey-Wilson

The colonization history of Olea europaea L. in Macaronesia based on internal transcribed spacer 1 (ITS-1) sequences, randomly amplified polymorphic DNAs (RAPD), and intersimple sequence repeats (ISSR)

Phylogenetic relationships in the Olea europaea complex and the phylogeography of 24 populations of the Macaronesian olive (O. europaea ssp. cerasiformis) were assessed by using three molecular markers: nuclear ribosomal internal transcribed spacer 1 (ITS-1) sequences, randomly amplified polymorphic DNAs (RAPD), and intersimple sequence repeats (ISSR). Parsimony analysis of the ITS-1 sequences and Neighbour-joining (NJ) analyses of RAPD and ISSR banding variation revealed four major lineages in the O. europaea complex: (1) ssp. cuspidata; (2) ssp. cerasiformis from Madeira; (3) ssp. laperrinei; and (4) ssp. cerasiformis from the Canary Islands plus ssp. europaea. These results provide unequ…

A comparative analysis of genetic variation in rootstocks and scions of old olive trees – a window into the history of olive cultivation practices and past genetic variation

Background Past clonal propagation of olive trees is intimately linked to grafting. However, evidence on grafting in ancient trees is scarce, and not much is known about the source of plant material used for rootstocks. Here, the Simple Sequence Repeat (SSR) marker technique was used to study genetic diversity of rootstocks and scions in ancient olive trees from the Levant and its implications for past cultivation of olives. Leaf samples were collected from tree canopies (scions) and shoots growing from the trunk base (suckers). A total of 310 trees were sampled in 32 groves and analyzed with 14 SSR markers. Results In 82.7% of the trees in which both scion and suckers could be genotyped, t…

Morphological and molecular evidence for interspecific hybridisation in the introduced African genusCrassocephalum(Asteraceae: Senecioneae) in Asia

Abstract The genus Crassocephalum in Asia, introduced there from Africa, was examined by extensive field work, herbarium studies, analyses of pollen and seed fertility, chromosome counts and ITS and trnL‐F sequencing. We found that Crassocephalum in Asia comprises two species and their interspecific hybrid. The two species are C. crepidioides (Benth.) S. Moore and C. rubens (Juss. ex Jacq.) S. Moore, of which the latter is a new record for Asia (north Thailand). The hybrid between these two species in north Thailand originated from a cross between C. crepidioides (2n = 40) as female and C. rubens (2n = 40) as male parent.

Phylogeographic evidence for the postglacial colonization of the North and Baltic Sea coasts from inland glacial refugia by Triglochin maritima L.

Abstract We investigated the geographical distribution of genetic variation in 67 individuals of Triglochin maritima from 38 localities across Europe using AFLP markers. Analysis of genetic variation resulted in the recognition of two major genetic groups. Apart from few geographical outliers, these are distributed (1) along the Atlantic coasts of Portugal, Spain and France and (2) in the North Sea area, the Baltic Sea area, at central European inland localities, the northern Adriatic Sea coast and the Mediterranean coast of southwest France. Considering possible range shifts of T. maritima in reaction to Quaternary climatic changes as deduced from the present-day northern temperature limit…

The phylogeny ofPapaver s. l. (Papaveraceae): Polyphyly or monophyly?

An RFLP analysis of the chloroplast genetrnK of 32 species of the generaPapaver, Roemeria, Stylomecon, andMeconopsis leads to the following conclusions: (1) AsianMeconopsis consists of two distinct clades and is paraphyletic in relation toPapaver, Roemeria, Stylomecon, and the W EuropeanMeconopsis cambrica. (2) Sister group relationships ofRoemeria toPapaver sect.Argemonidium and ofStylomecon toPapaver californicum are well-supported. (3)Meconopsis cambrica is nested withinPapaver (incl.Roemeria andStylomecon). The consideration of morphology, geographical distribution and ecology leads to the conclusion thatM. cambrica is best regarded as a member ofMeconopsis, and thatPapaver arose polyph…

Chloroplast DNA and isozyme analysis of the progenitor-derivative species relationship between Senecio nebrodensis and S. viscosus (Asteraceae )

A comparison of chloroplast DNA (cpDNA) and isozyme diversity in Senecio nebrodensis L., a species restricted to certain mountain ranges in Spain, and S. viscosus L., a widespread ruderal species in Europe, revealed that S. viscosus possessed the more common of two cpDNA haplotypes resolved in S. nebrodensis and contained only a small subset of the alleles found in S. nebrodensis at enzyme coding loci. The two species shared one restriction length mutation and one site mutation in their cpDNA, which distinguished them from other European Senecio species examined previously. Taken overall, these results support the hypothesis that S. nebrodensis and S. viscosus are related as a progenitor-de…

Evolutionary significance of the invasion of introduced populations into the native range of Meconopsis cambrica

The long history of the deliberate or accidental and human-mediated dispersal of flowering plants has led to the introduction of foreign genotypes of many species into areas of Europe hitherto occupied by potentially distinct native populations. Studies of the genetic and evolutionary consequences of such changes are handicapped by the difficulty of identifying the surviving native populations of many species in the absence of clear morphological differences. We investigated the relationship between putative native and introduced populations of the herbaceous perennial Meconopsis cambrica (Papaveraceae), as the isolated native populations of this species can be identified by historical and …

Drought tolerance of three olive cultivars alternatively selected for rain fed or intensive cultivation

Abstract Physiological drought response was evaluated for two olive cultivars commonly grown under rain fed conditions (‘Souri’ and ‘Picual’) and another selected for intensive, irrigated cultivation (‘Barnea’). ‘Souri’ is a traditional local Israeli cultivar, ‘Picual’ originated in Spain and ‘Barnea’ is a modern Israeli cultivar. Trees in pots were alternatively provided well irrigated conditions (100% FC, field capacity) or allowed to dry, first to 33% FC and then to 10% FC. Under conditions of greatest water availability, the ‘Barnea’ cultivar had the highest stomatal conductance and net photosynthesis, significantly higher than that found in ‘Souri’. Stomatal conductance and leaf water …

Walter Zimmermann and the Growth of Phylogenetic Theory

seminal theoretical paper, "Arbeitsweise der botanischen Phylogenetik und anderer Gruppierungswissenschaften" ("Methods of botanical phylogenetics and other grouping sciences"), first published in 1931. In this paper, Zimmermann clearly ex? pressed many of the underlying principles of phylogenetic systematics?ideas that were later taken up by Hennig and formed the core of his theory. Zimmermann also focused attention on several basic meth?

The phylogenetic relationships and evolution of the Canarian laurel forest endemicIxanthus viscosus (Aiton) Griseb. (Gentianaceae): Evidence frommatK and ITS sequences, and floral morphology and anatomy

The phylogenetic relationships of the Canarian laurel forest endemicIxanthus viscosus (Aiton) Griseb. (Gentianaceae) are investigated through a cladistic analysis of sequence variation of parts of the chloroplast genematK and the ITS region of nuclear ribosomal DNA. The floral anatomical and morphological characteristics ofIxanthus based on paraffin sections and SEM studies are also discussed. In the molecular analysesIxanthus is part of a clade of mostly temperate Erythraeinae and Chironiinae sensu Gilg, in which it is sister to the mostly Mediterranean annualBlackstonia. This relationship is supported by farreaching similarities in flower morphology and anatomy, and the shared possession …

Evolution of Leaf Morphogenesis: Evidence from Developmental and Phylogenetic Data in Papaveraceae

Variation of leaf morphology in Papaveraceae s.l. (including Fumariaceae and Pteridophyllum) has previously been shown to be related to developmental differences in the direction of segmentation and in blade‐petiole differentiation. Based on ontogenetic comparisons, we here distinguish polyternate, acropetal, basipetal‐pedate, basipetal‐pinnate, and divergent modes of dissection. In addition, undissected leaves occur in some taxa. Dissection modes can be grouped in two classes on the basis of blade‐petiole differentiation. Mapping of these morphogenetic traits on an independently obtained phylogenetic reconstruction reveals a high degree of homoplasy, indicating multiple evolutionary parall…

Which changes are needed to render all genera of the German lora monophyletic?

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Linnaeus's folly – phylogeny, evolution and classification ofSedum(Crassulaceae) and Crassulaceae subfamily Sempervivoideae

Stammesgeschichte und Systematik der Bakterien, Archaeen, „Pilze“, Pflanzen und anderer photoautotropher Eukaryoten

Nach der Entstehung des Lebens (s. Box 24.1) hat eine evolutionare Differenzierung der Lebewesen in drei Hauptgruppen stattgefunden, die meist als Reiche (Regnum/Regna) oder Domanen klassifiziert werden. Dies sind die Archaea, Bacteria und Eukarya. Gegenstand der Botanik und damit auch dieses Lehrbuchs sind Pflanzen sowie solche algenahnlichen photoautotrophen Organismen, die durch endocytobiotische Aufnahme von eukaryotischen Grun- oder Rotalgen entstanden sind. Diese Gruppen finden sich im Stammbaum des Lebens an unterschiedlichen Stellen (s. Abb. 24.1).

Genetic basis of speed of development in Senecio vulgaris L var. vulgaris, S. vulgaris ssp. denticulatus (O.F. Muell.) P.D. Sell, and Senecio vernalis Waldst. & Kit.

The genetic basis of differences in speed of development from germination to first bud formation was investigated in Senecio vulgaris var. vulgaris and S. vulgaris ssp. denticulatus, and also in S. vernalis sampled from Israel and Germany. In the case of S. vulgaris, F2 segregation analysis and the recovery of very late and very early lines from extreme F2 phenotypes showed that differences can be explained by a single major gene model, whereas segregation data from F2 and backcross progenies in S. vernalis are not incompatible with a digenic model of inheritance. Senecio vernalis from Israel and S. vulgaris var. vulgaris reached the different developmental stages in a substantially shorter…

Two new combinations inAdenostyles (Asteraceae, Senecioneae), a conspectus of the genus and key to its species and subspecies

Abstract Dillenberger M. S. & Kadereit J. W.: Two new combinations in Adenostyles (Asteraceae, Senecioneae), a conspectus of the genus and key to its species and subspecies. — Willdenowia 42: 57-61. June 2012. — Online ISSN 1868-6397; © 2012 BGBM Berlin-Dahlem. Stable URL: http://dx.doi.org/10.3372/wi.42.42105 Based on a molecular phylogeny (using ITS, ETS and the rpl32-trnL, psbA-trnH and ndhF-rpl32 plastid spacer sequences) of all species and subspecies of the European genus Adenostyles currently recognised, and supported by the morphology of the leaf margin, we transfer two subspecies of A. alliariae to A. alpina and provide the new combinations A. alpina subsp. macrocephala and A. alpin…

Molecular fingerprinting evidence (ISSR, Inter-Simple Sequence Repeats) for a wild status of Olea europaea L. (Oleaceae) in the Eurosiberian North of the Iberian Peninsula

Summary Although the olive tree ( Olea europaea L.) is one of the best indicators of the Mediterranean region, there exist some isolated populations of this species in the Eurosiberian North of the Iberian Peninsula. To elucidate the wild or feral status of these populations, we studied a total of 46 populations of the Olea europaea complex. mat K sequences and Inter-Simple Sequence Repeats (ISSR) variation were analysed to infer relationships among the six subspecies of the Olea europaea complex and within O. europaea subsp. europaea. Based on mat K. sequences, populations of sspp. europaea (Mediterranean basin), cerasiformis (Madeira), guanchica (Canary Islands), maroccana (Agadir Mountai…

Phylogenetic relationships, evolutionary origin, taxonomic status, and genetic structure of the endangered local Lower Elbe river (Germany) endemic Oenanthe conioides (Nolte ex Rchb.f.) Lange (Apiaceae): ITS and AFLP evidence

Abstract Oenanthe conioides is an endangered local endemic of the Lower Elbe river region in Germany where it is found in areas with freshwater tides. Internal transcribed spacer (ITS) and amplified fragment length polymorphism (AFLP) markers were used to investigate its phylogenetic relationships, evolutionary origin, taxonomic status, and genetic structure. Oe. conioides is most closely related to Oe. aquatica and Oe. fluviatilis, and cannot be distinguished from Oe. aquatica by ITS sequence variation. Oe. aquatica is found mainly in standing or slow-flowing freshwater habitats. The AFLP analysis indicated that Oe. conioides may constitute a monophyletic lineage nested within Oe. aquatica…

The phylogeny and biogeography of Gentiana L. sect. Ciminalis (Adans.) Dumort.: A historical interpretation of distribution ranges in the European high mountains

Abstract Gentiana sect. Ciminalis consists of seven mostly ecologically or geographically vicariant and closely related species which are distributed throughout the South and Central European high mountains. The analysis of a RAPD data set and trn L-intron and ITS sequences resulted in slightly different phylogenetic hypotheses. In the preferred hypothesis the group consists of two completely resolved main lineages: 1) G. clusii and G. alpina. 2) G. dinarica, G. acaulis, G. ligustica, G. angustifolia and G. occidentalis. The most important conclusions we have drawn from this phylogenetic hypothesis and from the observed patterns of molecular variation are: 1) The calcifuge ecology of G. aca…

Parallel evolution of flower reduction in two alpineSoldanellaspecies (Primulaceae)

The European endemic Soldanella has traditionally been divided into two morphologically well-defined sections. Section Tubiflores contains two species growing in high-elevation habitats, whereas most of the 14 species of section Soldanella inhabit montane forests. Section Tubiflores has a reduced floral morphology compared with section Soldanella. A previous phylogenetic study based on internal transcribed spacer (ITS) and AFLP data has revealed that, although the genus Soldanella itself is monophyletic, both sections are paraphyletic. Soldanella alpina (section Soldanella) forms a clade with S. minima and S. pusilla (section Tubiflores), and the grouping of S. alpina with S. pusilla has be…

Classification of Primula sect. Auricula (Primulaceae) based on two molecular data sets (ITS, AFLPs), morphology and geographical distribution

On the basis of the study of c. 1100 herbarium specimens, field observations and molecular (ITS, AFLPs) evidence, Primula sect. Auricula is classified into two subsections, 25 species and six subspecies. Primula auricula L. ssp. widmerae (Pax) L. B. Zhang stat. nov., P. auricula Linn. ssp. tatriaca L. B. Zhang ssp. nov., and P. latifolia Lapeyr. ssp. cynoglossifolia (Widmer) L. B. Zhang stat. nov. are newly described or combined; P. auricula s.l. is divided into two species: P. auricula Linn. and P. balbisii Lehm., representing the northern and the southern populations of P. auricula s.l. resolved in the molecular study, respectively; P. cottia Widmer and P. balbisii Lehm. are recovered fro…

Hybridization, ecogeographical displacement and the emergence of new lineages – A genotyping‐by‐sequencing and ecological niche and species distribution modelling study of Sempervivum tectorum L. (Houseleek)

Ecogeographical displacement of homoploid hybrid lineages from their parents is well documented and considered an important mechanism to achieve reproductive isolation. In this study, we investigated the origin of the flowering plant species Sempervivum tectorum in the Massif Central (France) through homoploid hybridization between lineages of the species from the Rhine Gorge area (Germany) and the Pyrenees (France). We used genotyping-by-sequencing genetic data as evidence for the hybrid origin of the Massif Central lineage, and WorldClim climatic data and soil pH and soil temperature data collected by us for ecological niche and species distribution modelling. We could show that the Massi…

Quaternary diversification in European alpine plants: pattern and process

Molecular clock approaches applied previously to European alpine plants suggest that Primula sect. Auricula , Gentiana sect. Ciminalis and Soldanella diversified at the beginning of the Quaternary or well within this period, whereas Globularia had already started diversifying in the (Late–)Tertiary. In the first part of this paper we present evidence that, in contrast to Globularia and Soldanella , the branching patterns of the molecular internal transcribed spacer phylogenies of both Primula and Gentiana are incompatible with a constant–rates birth–death model. In both of these last two taxa, speciation probably decreased through Quaternary times, perhaps because of some niche–filling pro…

Methoden der Systematik

Das Ziel der Systematik besteht darin, die enorme Vielfalt von Organismen unterschiedlichster Form und Lebensweise zu ordnen. Dies erfordert die Erkennung von Arten und deren Zusammenfassung zu systematischen Gruppen hoherer Rangstufe (Gattungen, Familien usw.). Zu den Aufgaben der Systematik gehort es auch, Arten und hohere systematische Gruppen zu beschreiben und zu benennen und in Form von Bestimmungsschlusseln Moglichkeiten fur deren Identifizierung zur Verfugung zu stellen. Seitdem bekannt ist, dass alle heute lebenden Organismen von einem gemeinsamen Vorfahren abstammen (s. Evolution, Kap. 22) ist die Systematik darum bemuht, die naturlichen Verwandtschaftsverhaltnisse zwischen Organi…

Is Welsh Poppy,Meconopsis cambrica(L.) Vig. (Papaveraceae), truly aMeconopsis?

AbstractSince the nineteenth century, the western European endemic Meconopsis cambrica has been regarded as the only European representative of the genus Meconopsis Vig. This genus, which is otherwise restricted to the Himalayas, differs from Papaver in having a style rather than a stigmatic disc. A phylogenetic reconstruction using 65 internal transcribed spacer sequences of 62 taxa of Old World Papaveroideae and three outgroup taxa shows that M. cambrica is not the closest relative of the remainder of Meconopsis but rather the closest sampled relative of Papaver s.str. This is consistent with morphological evidence which suggests that the style evolved independently in M. cambrica from a …

Ecological implications of Cousinia Cass. (Asteraceae) persistence through the last two glacial–interglacial cycles in the continental Middle East for the Irano-Turanian flora

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Typification and synonymization inPrimulasect.Auricula(Primulaceae)

Supplementing a recent systematic study of Primula sect. Auricula, we here lectotypify or neotypify 27 names applicable to taxa in this section and the name of one related genus. These are P. auricula var. widmerae Pax, P. balbisii Lehm., P. ciliata Moretti, P. cottia Widmer, P. crenata Lam., P. oenensis Thomas ex Gremli, P. glaucescens Moretti, P. glutinosa Wulfen, P. integrifolia L., P. latifolia var. cynoglossifolia Widmer, P. laevigata Duby ex Rchb., P. longobarda Parta, P. lutea Vill., P. marginata Curtis, P. microcalyx Lehm., P. minima L., P. parlatorii Porta ex Caruel, P. villosa Wulfen, P. villosa var. daonensis Leyb., P. viscosa Vill., P. [unranked] Brevihracteae Widmer, P. [unrank…

A conspectus of Tephroseris (Asteraceae: Senecioneae) in Europe outside Russia and notes on the decline of the genus

Tephroseris is generally considered a difficult genus. Based on the examination of extensive herbarium material and considering the existing literature, we recognize seven species in Europe outside Russia. These are T. palustris, T. integrifolia with subsp. integrifolia, subsp. aurantiaca, subsp. capitata, subsp. maritima, subsp. serpentini and subsp. “tundricola”, T. balbisiana, T. crispa, T. helenitis, T. longifolia and T. papposa. Phylogenetic analysis of ITS and ETS sequences showed that these species fall into three lineages. These are: (1) T. palustris, clearly related to Arctic species of the genus; (2) T. integrifolia; and (3) the remaining species. Molecular dating of the T. integr…

Molecules and Morphology, Phylogenetics and Genetics

Various explanations can be offered for the incongruence between phylogenetic hypotheses resulting from morphological and molecular data sets. Of these, the possibility that incongruence may result from the mutation of major morphogenetic genes leading to dramatic morphological divergence unaccompanied by equivalent change of the phylogenetic marker molecule(s) used is discussed in detail. As evidence for this hypothesis, several examples for such incongruence are surveyed. It seems possible that in many cases the genetic basis of the morphological characters responsible for the incongruence found may be simple, and that the genes involved may be homologous to genes known from mutant system…

A revision of Papaver sect. Meconidium

The biennial Papaver L. sect. Meconidium Spach (Papaveraceae) is considered to contain four species: P. libanoticum Boiss. with subspp libanoticum and polychaetum (Schott & Kotschy ex Boiss.) Kadereit comb. et stat. nov., P. armeniacum (L.)DC. with subspp armeniacum, microstigmum (Boiss.) Kadereit comb. et stat. nov. and pilgerianum (Fedde) Kadereit comb. et stat. nov., P. curviscapum Nab., and P. persicum Lindl. with subspp persicum, tauricolum (Boiss.) Kadereit comb. et stat. nov. and microcarpum (Boiss.) Kadereit comb. et stat. nov. A key to species and subspecies is given, capsules and leaves of all taxa are illustrated, and distribution maps are provided.

The evolution of substrate differentiation inMinuartiaseriesLaricifoliae(Caryophyllaceae) in the European Alps: In situ origin or repeated colonization?

Premise of the study Substrate specialization is often considered an important factor in evolutionary diversification. A classic example of divergence related to different substrate types is the dichotomy between calcicole and calcifuge plants on calcareous and siliceous substrates as found in the European Alps. When closely related species with contrasting substrate preferences are found in the same area, it is generally hypothesized that they diverged where they now occur. However, it is possible that Alpine edaphic diversity instead allows the coexistence of related species whose edaphic differentiation took place deeper in the phylogeny, in some other part of the range of their clades. …

Frequent colonization and little in situ speciation in Senecio in the tropical alpine-like islands of eastern Africa.

Premise of the study Floras of continental habitat islands, like those of islands, originate mostly through colonization, which can be followed by in situ speciation. We here address the question of the relative importance of colonization and in situ diversification in the high-altitude areas of the eastern African high mountains, the tropical Afroalpine Region, using the most species-rich genus in the region, Senecio, as an example. Methods We expanded earlier Senecioneae phylogenies by adding more tropical African species and analyzed our phylogenetic tree biogeographically. Key results Senecio contains at least five clades with tropical African species, all of them containing tropical af…

REVISION OF THE MEDITERRANEAN AND SOUTHERN AFRICAN TRIGLOCHIN BULBOSA COMPLEX (JUNCAGINACEAE)

The Triglochin bulbosa complex (Juncaginaceae) from the Mediterranean region and Africa is revised. One new species, Triglochin buchenaui Köcke, Mering & Kadereit, and two new subspecies, Triglochin bulbosa subsp. calcicola Mering, Köcke & Kadereit and Triglochin bulbosa subsp. quarcicola Mering, Köcke & Kadereit, are described from South Africa. The only two Mediterranean taxa in the complex (Triglochin barrelieri, T. laxiflora) are elevated to species rank. Altogether seven species and four subspecies are recognised: Triglochin barrelieri, T. buchenaui, T. bulbosa subsp. bulbosa, T. bulbosa subsp. calcicola, T. bulbosa subsp. quarcicola, T. bulbosa subsp. tenuifolia, T. compac…

Causes of the genetic architecture of south-west European high mountain disjuncts

Background: Postglacial climatic warming in south-western Europe and the retreat of cold-adapted species into higher elevations, starting in the Sierra Nevada and proceeding northwards to the Pyrenees and Alps, should have resulted in a pattern of ‘successive vicariance’. Alternatively, long-distance dispersal might explain the extant distribution pattern of mountain species in this region. Aims: Here, we report an investigation of two alpine plants, Saxifraga oppositifolia and S. stellaris, which co-occur in the Sierra Nevada, the Pyrenees, and the south-western Alps/Massif Central. Our aim was to distinguish between (successive) vicariance and long-distance dispersal as alternative explan…

Chloroplast DNA and isozyme evidence on the evolution ofSenecio vulgaris (Asteraceae)

Chloroplast DNA (cpDNA) and isozyme variation were analyzed over a range of populations of two infraspecific taxa of the tetraploidSenecio vulgaris. The isozyme data were supportive of the hypothesis that the weedy and cosmopolitanS. vulgaris var.vulgaris is an evolutionary derivative ofS. vulgaris subsp.denticulatus from the coasts of W Europe and montane altitudes in S Spain and Sicily. The two taxa exhibited a very high genetic identity with subsp.denticulatus containing slightly more isozyme diversity than was found in var.vulgaris. — Three cpDNA haplotypes (A, B, C) already known from other Mediterranean diploid species ofSenecio were resolved in var.vulgaris, and an additional fourth …

Grundlagen der Vererbung

Genetische Variation entsteht durch Mutation, aber auch durch die Durchmischung des Erbguts unterschiedlicher Individuen. Dieser als Rekombination bezeichnete Prozess ist bei eukaryotischen Organismen an die sexuelle Fortpflanzung gebunden. Die Rekombination des elterlichen Erbguts wird einerseits durch die Zufalligkeit der Verschmelzung von Keimzellen (Syngamie) und andererseits durch die Vorgange der meiotischen Zellteilung bei der Entstehung der Gameten der nachsten Generation bewirkt. Die Erzeugung genetischer Variation ist eine, aber moglicherweise nicht die einzige, wichtige Funktion der Sexualitat.

Parallel bursts of recent and rapid radiation in the Mediterranean and Eritreo-Arabian biodiversity hotspots as revealed byGlobulariaandCampylanthus(Plantaginaceae)

Evolution of gynoecium morphology in Old World Papaveroideae: a combined phylogenetic/ontogenetic approach.

PREMISE OF THE STUDY: The correct assessment of homology is an important prerequisite for reconstructing phylogenetic relationships and character evolution. Old World Papaveroideae (Papaver, Meconopsis, Roemeria, Stylomecon) show substantial diversity in gynoecium and capsule morphology. In particular, capsules can have distinct styles (Meconopsis p.p., Stylomecon) or a sessile stigmatic disc (Papaver). Molecular phylogenetic analyses of Old World Papaveroideae had shown that neither taxa with styles nor those with stigmatic discs represent monophyletic lineages. We here investigate whether either styles or stigmatic discs have arisen repeatedly during the diversification of Old World Papav…

An AFLP clock for the absolute dating of shallow-time evolutionary history based on the intraspecific divergence of southwestern European alpine plant species.

The dating of recent events in the history of organisms needs divergence rates based on molecular fingerprint markers. Here, we used amplified fragment length polymorphisms (AFLPs) of three distantly related alpine plant species co-occurring in the Spanish Sierra Nevada, the Pyrenees and the southwestern Alps/Massif Central to establish divergence rates. Within each of these species (Gentiana alpina, Kernera saxatilis and Silene rupestris), we found that the degree of AFLP divergence (D(N72)) between mountain phylogroups was significantly correlated with their time of divergence (as inferred from palaeoclimatic/palynological data), indicating constant AFLP divergence rates. As these rates d…

Reciprocal hybridization at different times between Senecio flavus and Senecio glaucus gave rise to two polyploid species in north Africa and south‐west Asia

Summary • The analysis of hybrid plant taxa using molecular methods has considerably extended understanding of possible pathways of hybrid evolution. • Here, we investigated the origin of the tetraploid Senecio mohavensis ssp. breviflorus and the hexaploid Senecio hoggariensis by sequencing of nuclear and chloroplast DNA, and by analysis of the distribution of taxon-specific amplified fragment length polymorphism (AFLP) fragments. • Both taxa originated from hybridization between the diploid Senecio flavus and Senecio glaucus. Whereas S. glaucus was the female parent in the origin of S. mohavensis ssp. breviflorus, S. flavus was the female parent in the origin of S. hoggariensis. • The dist…

The geography of hybrid speciation in plants

The identity and geographical distribution of Jacobaea vulgaris subsp. gotlandica , supposedly endemic to Gotland and Öland (Sweden)-the importance of multiple intraspecific samples

The identity of Jacobaea vulgaris Gaertn. subsp. gotlandica (Neum.) B. Nord., hitherto considered an endemic to the Swedish islands Oland and Gotland, was investigated using nrITS and trnT-trnL intergenic spacer sequences and a broad sample of species and accessions particularly of J. vulgaris. We found that J. vulgaris subsp. gotlandica is a distinct evolutionary lineage more widespread in Europe than previously thought and may be a southeastern European steppe floristic element with outliers on the large Baltic islands. Our entirely unexpected findings underline the need of multiple intraspecific sampling in species-level phylogenetic studies.

Identity of the Calcarata species complex in Viola sect. Melanium (Violaceae)

The Calcarata species complex in Viola sect. Melanium (Violaceae) is a group of species from Italy and neighbouring islands. The complex is of considerable evolutionary interest because several hypotheses about hybrid speciation within the group have been previously proposed. Because the Calcarata complex is not well characterized morphologically, we used 142 samples representing 92 (of c. 120) species of V. sect. Melanium plus three outgroup species. Nuclear ITS and ETS and plastid trnS–trnG intergenic spacer sequences were analysed to test the monophyly of the Calcarata complex and to infer relationships among the constituent species. Both nuclear and plastid sequences resulted in very li…

Maximum polyphyly: Multiple origins and delimitation with plesiomorphic characters require a new circumscription of Minuartia (Caryophyllaceae)

Testing the potential significance of different scion/rootstock genotype combinations on the ecology of old cultivated olive trees in the southeast Mediterranean area

Background A previous multi-locus lineage (MLL) analysis of SSR-microsatellite data of old olive trees in the southeast Mediterranean area had shown the predominance of the Souri cultivar (MLL1) among grafted trees. The MLL analysis had also identified an MLL (MLL7) that was more common among rootstocks than other MLLs. We here present a comparison of the MLL combinations MLL1 (scion)/MLL7 (rootstock) and MLL1/MLL1 in order to investigate the possible influence of rootstock on scion phenotype. Results A linear regression analysis demonstrated that the abundance of MLL1/MLL7 trees decreases and of MLL1/MLL1 trees increases along a gradient of increasing aridity. Hypothesizing that grafting o…

The phylogeny of (Gentianaceae) and its colonization of the southern hemisphere as revealed by nuclear and chloroplast DNA sequence variation

Abstract The generic circumscription and infrageneric phylogeny of Gentianella was analysed based on matK and ITS sequence variation. Our results suggested that Gentianella is polyphyletic and should be limited to species with only one nectary per petal lobe. Gentianella in such a circumscription is most closely related to one part of a highly polyphyletic Swertia. within uninectariate Gentianella two major groups could be recognized: 1) northern hemispheric species with vascularized fimbriae at the base of the corolla lobes, and 2) northern hemispheric, South American, and Austrlia/New Zealand species without vascularized fimbriae. When fimbriae are present in this latter group, they are n…

Local old olive landrace varieties in Israel—Valuable plant genetic resources in olive cultivation

It is assumed that the cultivation of olive trees started in the East Mediterranean in the third millennia BCE. Throughout history and until recently, successful olive clones were maintained vegetatively and were grafted either on seedlings or on spheroblasts removed from the base of the trunk. It therefore can be hypothesized that local old olive trees, older than 500 years might represent an ancient gene pool. Modern terminology categorizes local genotypes into four main variety groups. However, in traditional terminology these were further differentiated into 27 cultivars according to their phenotypic traits. This genetic diversity, along with the fact that olive trees have been cultivat…

The influence of sea currents, past disruption of gene flow and species biology on the phylogeographical structure of coastal flowering plants

Aim We investigate the geographical genetic structure of two coastal plant species, Cakile maritima Scop. (Brassicaceae) and Eryngium maritimum L. (Apiaceae), through three sea straits and along one continuous stretch of coast using amplified fragment length polymorphisms (AFLPs). The two species have a similar ecology in that they grow in sandy habitats, but differ in life-form (annual vs. perennial) and dispersability of seeds by sea water as inferred from floating experiments. The sea straits differ in their geological history and their modern current systems. The primary goal of our study was to test the hypothesis that sea straits have an influence on the geographical patterns of genet…

A note on leaf venation and the circumscription of Tephroseris (Asteraceae–Senecioneae)

Molecular phylogenetic results had shown that Tephroseris, a genus differentiated from its closest relatives by its pinnately veined leaves, also contains species with palmate leaf venation. This had led to the conclusion that leaf venation is a homoplasious character in the lineage containing Tephroseris. We here take a closer look at leaf venation in Tephroseris and other species of Asteraceae–Senecioneae. We found that leaf venation in Tephroseris is distinct from leaf venation in Jacobaea vulgaris and Senecio vulgaris, which both have pinnately veined leaves as typical for most species of their respective genera, and that description of the leaves of Tephroseris as pinnately veined is i…

The origin of the serpentine endemic Minuartia laricifolia subsp. ophiolitica by vicariance and competitive exclusion.

Serpentine soils harbour a unique flora that is rich in endemics. We examined the evolution of serpentine endemism in Minuartia laricifolia, which has two ecologically distinct subspecies with disjunct distributions: subsp. laricifolia on siliceous rocks in the western Alps and eastern Pyrenees and subsp. ophiolitica on serpentine in the northern Apennines. We analysed AFLPs and chloroplast sequences from 30 populations to examine their relationships and how their current distributions and ecologies were influenced by Quaternary climatic changes. Minuartia laricifolia was divided into four groups with a BAPS cluster analysis of the AFLP data, one group consisted only of subsp. ophiolitica, …

Repeat intercontinental dispersal and Pleistocene speciation in disjunct Mediterranean and desert Senecio (Asteraceae).

To explore the biogeographic history of Mediterranean/arid plant disjunctions, Old and New World Senecio sect. Senecio were analyzed phylogenetically using nuclear ribosomal DNA sequences (ITS). A clade corresponding to sect. Senecio was strongly supported. Area optimization indicated this clade to be of southern African origin. The Mediterranean and southern African floras were not distinguishable as sources of the main New World lineage, estimated to have become established during the middle Pliocene. Another previously suspected recent dispersal to the New World from the Mediterranean was confirmed for the recently recognized disjunction in S. mohavensis. The loss of suitable land connec…

Combining palaeodistribution modelling and phylogeographical approaches for identifying glacial refugia in AlpinePrimula

Aim We investigated the late Quaternary history of two closely related and partly sympatric species of Primula from the south-western European Alps, P. marginata Curtis and P. latifolia Lapeyr., by combining phylogeographical and palaeodistribution modelling approaches. In particular, we were interested in whether the two approaches were congruent and identified the same glacial refugia. Location South-western European Alps. Methods For the phylogeographical analysis we included 352 individuals from 28 populations of P. marginata and 172 individuals from 15 populations of P. latifolia and used amplified fragment length polymorphisms (AFLPs). For palaeodistribution modelling, species distrib…

Differential cycles of range contraction and expansion in European high mountain plants during the Late Quaternary: insights from Pritzelago alpina (L.) O. Kuntze (Brassicaceae).

Nuclear DNA sequence variation of the internal transcribed spacer (ITS) and amplified fragment length polymorphisms (AFLPs) were used to illuminate the evolutionary history of Pritzelago alpina, a herbaceous perennial of (sub)alpine to nival habitats of the European high mountains. Maximum likelihood analysis of ITS sequences of P. alpina, Hornungia petraea and Hymenolobus procumbens (the 'Pritzelago alliance') resolved P. alpina and H. petraea as sister taxa. ITS divergence estimates support an origin for P. alpina in the Late Tertiary, while intraspecific diversification started in the Late Quaternary (0.4-0.9 million years ago). AFLP analysis of 76 individuals of P. alpina, representing …

Long‐distance dispersal vs vicariance: the origin and genetic diversity of alpine plants in the Spanish Sierra Nevada

Here, we investigated the origin and genetic diversity of four alpine plant species co-occurring in the Spanish Sierra Nevada and other high mountains in south-western Europe by analysis of amplified fragment length polymorphisms (AFLPs). In Kernera saxatilis, Silene rupestris and Gentiana alpina we found intraspecific phylogroups corresponding to mountain regions as predicted by the vicariance hypothesis. Moreover, genetic distances between Sierra Nevada and Pyrenees populations were always higher than those between populations from the Pyrenees and the south-western Alps/Massif Central. This suggests successive disruption of gene exchange between mountain ranges as postglacial climatic wa…

Three intercontinental disjunctions in Papaveraceae subfamily Chelidonioideae: evidence from chloroplast DNA

An RFLP analysis of the chloroplast genome of Papaveraceae subfam. Chelidonioideae resulted in one most parsimonious tree consisting of three monophyletic groups. Each group contains intercontinental disjunct taxa. Whereas Eomecon/Sanguinaria and Stylophorum are examples for the well known E Asian — eastern N American disjunction, the E Asian -C and S American disjunction in Macleaya/Bocconia is rare. The genus Stylophorum in this analysis is paraphyletic. The N American Stylophorum diphyllum is sister group to the Old World members of Stylophorum and Chelidonium. All American representatives of the subfamily possess distinctive morphological features. This might be the result of Tertiary a…

Terrestrial growth and marine dispersal? Comparative phylogeography of five coastal plant species at a European scale

The phylogeography of five flowering plant species (Cakile maritima, Eryngium maritimum, Salsola kali, Halimione portulacoides, Crithmum maritimum) widespread along the European coasts was investigated across their entireEuropean range using AFLP evidence. Both similarities and dissimilarities were found. All species contain a distinct Black Sea/Aegean Sea cluster, and all except E. maritimum contain a distinct Adriatic Sea cluster or group of genetically very similar clusters. All species except Cr. maritimum contain a distinct Atlantic Ocean/North Sea/Baltic Sea cluster clearly separate from the Mediterranean material. In Ca. maritima a distinct Baltic Sea subcluster was found. In the wes…

Systematics, Phylogeny, and Evolution of Papaver californicum and Stylomecon heterophylla (Papaveraceae)

Abstract We present a detailed comparison of Papaver californicum and Stylomecon heterophylla, which earlier were found to be sister species and most closely related to Meconopsis cambrica + Papaver s.str. from western Eurasia. The two species of winter annuals differ mainly in the shape of their distal cauline leaves, coloration of petals and staminal filaments, and most notably morphology of the gynoecium and capsule, with Papaver californicum having a sessile stigmatic disc and Stylomecon heterophylla having a distinct style. They were earlier found to differ in ploidy, with chromosome numbers of 2n  =  28 (Papaver californicum) and 2n  =  56 (Stylomecon heterophylla). Mapped distributio…

The phylogeny of the European high mountain genus Adenostyles (Asteraceae-Senecioneae) reveals that edaphic shifts coincide with dispersal events.

UNLABELLED PREMISE OF THE STUDY Heterogeneity of edaphic conditions plays a large role in driving the diversification of many plant groups. In the Alps and other European high mountains, many closely related calcicole and calcifuge plant taxa exist. To better understand patterns and processes of edaphic differentiation, the phylogeny of the edaphically variable genus Adenostyles was studied. The genus contains three species, of which A. alpina has five subspecies. Each species and subspecies is largely confined to either calcareous or noncalcareous substrates. • METHODS We analyzed the phylogeny of Adenostyles using DNA sequences of nrITS, nrETS, nuclear chalcone synthase, and three plastid…

Spatial and Temporal Patterns in the Evolution of the Flora of the European Alpine System

This paper presents a perspective of how phylogenetic and phylogeographic hypotheses, based on nuclear DNA sequence variation (ITS) or amplified fragment length polymorphisms (AFLPs), can provide insights into the origin and evolution of the European high mountain flora. We focus on a diversity of unrelated herbaceous plant taxa that are broadly co-distributed across the European Alpine System, representing different taxonomic levels, and having either Mediterranean or Asian affinities (i.e., Anthyllis montana, Pritzelago alpina, Globularia vs. Soldanella, and Primula sect. Auricula). Our observations highlight that all taxa investigated began to diversify at the beginning of the Pleistocen…

Data from: Phylogeography of a Tertiary relict plant, Meconopsis cambrica (Papaveraceae), implies the existence of northern refugia for a temperate herb

The perennial herb Meconopsis cambrica, a western European endemic, is the only European species of the otherwise Himalayan genus Meconopsis and has been interpreted as a Tertiary relict species. Using rbcL and ITS sequence variation, we date the split between M. cambrica and its sister clade Papaver s.str. to the Middle to Upper Miocene (12.8 My, 6.4-19.2 My HPD). Within M. cambrica, cpDNA sequence variation reveals the existence of two groups of populations with a comparable level of genetic variation: a northern group from Great Britain, the Massif Central, the western Pyrenees and the Iberian System, and a southern group from the central and eastern Pyrenees. Populations from the Cantab…

Data from: The origin of the serpentine endemic Minuartia laricifolia subsp. ophiolitica by vicariance and competitive exclusion

Serpentine soils harbour a unique flora that is rich in endemics. We examined the evolution of serpentine endemism in Minuartia laricifolia, which has two ecologically distinct subspecies with disjunct distributions: subsp. laricifolia on siliceous rocks in the western Alps and eastern Pyrenees and subsp. ophiolitica on serpentine in the northern Apennines. We analysed AFLPs and chloroplast sequences from 30 populations to examine their relationships and how their current distributions and ecologies were influenced by Quaternary climatic changes. Minuartia laricifolia was divided into four groups with a BAPS cluster analysis of the AFLP data, one group consisted only of subsp. ophiolitica, …